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The Evolution of Angiospern1id Conjectures and Pollen Queries

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The Evolution of Angiospern1id Conjectures and Pollen Queries The evolution of angiospern1id pollen characteristics: conjectures and queries G Vasanr!w, B S \'enkarachala & S. A J Pocock \·:.IS:.InIl1\, G .. \·enkataclub. H. S. Co: Pocock.~. A. .J. 1990. 111C e\'ollllion ofangiospermid pollen cluractcristlCS: conjecture, and querie.'. III Jain. K P. Co: Tiwari, R. S (ecls )-I-'roc Sl'lIIP '\'islas ill Ilie/iall fJa!a('o!Jolall)" l-'ala('o!Jolaliisl 38 UI 146. The migin and e\OIUllOll of dillcrent eXll1e la\'ers Ill' pah-nofossib is analysed in the light of accumulating ullrastruCtural dala. Semi·diagramll1:.tllc illustrations ba,ed on the published TE~I results of \'arious exil1t' l\'pes represel1tillg a no" ,eCliol1 of' e:-aincr and eXl3nt plant groups arc givcn I'or cas\' refcrelKe and cOll1prehCnslllll. ~ome "I' the imponalH pah'nological questions anel issues discussed in the present \york Me: imprecise use. to dcscribe the infra·tectum of pollen. Ill' the nexible term "granular" that of'ten leads to erronellUS ckri\ations and conclusiol1S; ontogenctic differences between the apparently similar complexh' ah-eolate columellate sexine tl'pes of g\'lllnosperms and :.Ingiosperms respecli\'elv; independelH e\'olution of columellar cOlllplexi!\' in unrf'laled taxa: role of ubiquitous "'hite lines in the exines of extinCt and extailt spores and pollen alld adaptive rather thall plwlogenetic significance 01' sacci in progyJl)nosperms, ~'mnosperms and angiosperms, Despite recognition of angiospermid pollen charaCteristics especiallv in tecralll' reticulate and colull1ellate pnllen of Tri:.tssi<: (Cornet, 7 J9 9, 19H~. 1989; Pocock Co: Vasanth\', 198R; Pocock, \'asamhy & VenKatachala. 19H8) the pre Cretaceous origin of angiosperm st ill remain~ an open quest ion. Key-words - Compara( i\ e anah·sis. Exine ulirastruc( lire, Angiosperm e\'olut ion. G \'ClSClI/II.~)'. Frel/ch II/,;Iilille uJ PUI/Cl!Cb"ITI', J() Sf. I,Ullis Slre<'!. PI3 No ,U. POl/c!lc/?('n:r 605 OOJ. II/aw 8, S l'eIlRaI(/c!>"la, Hir!J(/! SCi/II// 11/,;/1111/1' oJ 1'(/laeo!)oICiI/Y, 'ij {'I/II'ersi/.l' Hoad, Lllc!,>lIuu' 120 007, Il/uiC/. S A j. Poco c!,>, /-I.H.I., Cre,;/ol/. He. "on IG0. Cal/{/e/a. ri~T ~~<f;",~~~.~'f'i''1"~'" ~ ~ ~ ~ ~o ~ ;;\to 'lTf1r~, ttBo c;o ~ ~ fqN?[~~ ~'i[q ~ ~ ~ ~ ~~ Q(lfI<'1'1lcGqi :mq;if <f; mUll: Gl: <tit <liT lfllT%1 ~ ~ ~ ~Tc1 ~ ~ ~Vf ani-~ qrctI' ;j; fqf'l[?[ g<m <f; <tit 3!'!"f'll' <fTif q<: <f; 'If<crrrq'f Gl: mmf'm ~~ 'T~" ~ ~ ~ ~ '~J;m'~, ~ ~ ~ m 'l\T ~ fu;ir llit ~I ~mu-1ft if Q,I"lulf'qil> 'T'liT'< '1fll': ~ ~ ~, ~ ~n al'1I~<1.n"'~ldr 3f1~<1"1~I"'~llli ~~T ~ ~/~ ~ ~ ~ llit <liT mf\:ra' 1J1it1T; c;q' <f; <f; 'l"<IiTiiif~~fqf\r~;~~iifd''liT~<IiT~~;~~~~~~m<f;~Gl: ~ ~ ~ 'l<l3f'11~<1"11"'1, 3f'11~<141"'1 ~ 3f1~<141"'11l1 '~' ~ ~ ~ ~ wk tIDm <tit ,PIT <f; <f; I ~ (~, ~ 'lTf1r~, ~, 'lTf1r~ q~, ~ ~ ~ <Fi"T 1979, 1985, 1989; q 1988: 1988) <f; fd''liT ~ ~ ~ ~ ~ 'J:<I-m~iT ~'i[q ~ 'T~ "Ilfi1qil'l'"! if :J!T'floft;;jt <f; <'flll"i! <lit 'llRf<1T <f; tfrm <liT 31'l\T 'liT 0l,<1T %I "POLLEN characters are subject to parallelism, emphasise the need for detailed palynological convergence and possible reversal and the study of descriprions and critical analyses of exine fine structural derails of fossil pollen opens up characteristics of pre-Cretaceous palynofossils. possibilities of new sources of phylogenetiC Light microscopic studies and SEM observations evidence" (Davis & Heywood, 1963). This paper is a of various spore-pollen types have already amply follow-up of our recent contributions (Pocock & discussed and hypothesised the evolution of Vasanthy, 1988; Pocock, Vasanthy & Venkatachala, germinal apertural types through ages. The objective 1988; Vasanthy, Venkatachala & Pocock, 1988; of our present study was to trace the phylogenetiC Pocock, Vasanthy & Venkatachala, 1990) which relationships with the aid of published 131 \32 THl PALAE0I30TANIST ultrastructural details of palynofossils from hypothetically derived from "atectate" exines Palaeozoic through Cretaceous. For easy reference (Doyle, Van Campo & Lugardon, 1975; Walker & and comprehension we have included in this paper Skvarla, 1975). Crane (1985; table 9) while the semidiagrammatic illustrations of 56 TEM reviewing the occurrence of granular exine in pictures of exine types. Legends of these Text­ various plant groups (e.g. Archaeopteris, figures Include our comments, conjectures and CorystOspermales, Bennettitales, Gnetales and many queries chIefly pertaining to basic palynology, which angiosperms) accepted the hypothesis that granular may aid palynologists to appreciate the new pollen wall stratification is primitive within dimensions of palyno-phylogeny. angiosperms (cf Doyle, 1978). From an analysis of ultrastructural data of Ultrastructure of "granular" types of exines in palynofossll and pollen types we infer that: (i) the fossil and extant gymnospermous pollen has been polyplicate pollen morphotypes of Equisetosporites, the subject of many palynological papers. Foster and Ephedrzpites, Ephedra and Spathiphyllum (Aractae) Price (1981) described the Permian palynofossil are infrastructurally different, thus negating any Praecolpatites sinuosus (Text-fig. 3 A) as "incipiently phylogenetic relationship among them; (ii) pre­ alveolate" and even compared its "cavitate" or Cretaceous Equisetosporites chinLeanus and "granular" exoexine to the "granular" exine of Cornetipollis relicuLata are columellate, like certain Magnoliaceae. Zavada (1984) considered the Cretaceous angiosperms (cf. Pocock & Vasanthy, exine of Praecolpatites as a granular form. Should we 1988); (iii) prevalence of saccus in many non­ then infer that the terms "granular" and "inCipiently angiospermous plant groups and in primitive alveolate" are synonyms or equivalents? If so, the angiosperm Lactoridaceae is not a good character to exine of Middle Jurassic Corystosperm Pteruchus link angiosperms with gymnosperms; (iv) dubius (Zavada & Crepet, 1985; Text-fig 5 C) could columellar complexity might have risen be described as either granular or incipiently independently in extant Coniferae, Circumpolles, alveolate. The ultrastructure of another some Eocene fossils and many taxa of angiosperms; gymnospenTIid Mesozoic palynofossil Eucommiidites (v) the tripartite nexine of Equisetosporites (Doyle et aI, 1975; Text-fig. 1 A) reveals an chinLeanus with lamellate inter-bedded, mid-zone anastomosing granular infratectum (prelude to may be representative of a transitional evolutionary columellar evolutionl) Although Zavada (1984) stage, intermediate between lamellate described the infratectum of bisaccate pollen of gymnospermous and non-Iamellate angiospermous Triassic, Jurassic and Cretaceous as granular, the nexine; and (vi) as myriads of exine-types have been Cenomanian Granabivesiculites sp. cf. G. inchoatus imprecisely described as "granular" and "spongy", and the Albian vestigial saccate pollen (Text-fig. 3 B, one ought to be very discreet while grouping these C) look complexly columelliform (anastomosing types to draw phylogenetiC inferences. rods) rather than granular. Likewise the dispersed monosulcate pollen from the Cenomanian Dakota GRANULAR INFRATECTUM : FACTS AND Formation (Zavada & Dilcher, 1988) are apparently FALLACIES columellar or distinctly columelloid (e.g Text-fig. 5 D· F) but not granular. In pollen morphological descriptions, the term Amongst the extant gymnosperms, the exines of granular is used in its broadest sense: to describe Agathis aLba (Araucariaceae) and Cupressus the sexinal or tectaI surface sculpture (cf. Kremp, arizonica (Cupressaceae) have granular 1965, p. 61); the infra-tectal interstitial structure and ultrastructure (Van Campo & Lugardon, 1973). But in sometimes even the nexine. Van Campo and the latter (Text-fig. 3 D) the granular infratectum Lugardon (1973) defined granular structure as tends to be columelloid. Granular pollen sporopollenin organised into spherical grains, more development in Cunninghamia LanceoLata or less densely distributed under the tectum and (Taxodiaceae) has been investigated (Kurmann, generally discernible under the electron 1988) The pollen wall of Gnetales is generally microscope. The "synonyms" of granular classified under granular type (Text-fig. 1 B, C, D). infratectum-a network of rods of various size, The inter·crestal infratectum of Ephedra navadensis poorly developed columellae, columellae formed of (Cornet, 1985) and E. distachya (Van Campo & small granules, granular layer of endosexine and Lugardon, 1973) is columelloid unlike the others-have been discussed in their informative granuliform infra-tectum of arched summit of the review on granular exine. ridges. In WeLwitschia mirabiLis (Gullvag, 1966; "Granular" exines that occur both in Hesse, 1984), the interstitium is granulo-reticuloid gymnosperms and angiosperms, have been (tending to be columelloidl) The infratectum of the VASAJ'HHY et al.-[VOLUTION Of A!\lGIOSPERJv!Jl) POI.I.EN CHARACTERISTICS I )3 inter-spinular areas in Gnetum spp. (Gullvag, 1966; Mimosoideae (Guinet & Barth, 1967), anastomosed Zavada, 1984) tends to be columelloid unlike the granules orienting into columellae in Vigna crowded granules of the domed infra-spinular longifolia, Papilinoideae (Horvat & Stainier, 1980) interstitium. Some of the aforementioned examples and the columellae secondarily reversing to may be possibly or probably indicative of granulo­ granuliform infratectum accompanied by ross of columellar transitional stages of infratectum in non· footlayer in the Orchidaceae (Burns-Baloch & Hesse, angiosperms. 1988) are a few examples (Text-fig. 2) supportive of The prevalence
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